Fruit characters as a basis of fruit choice and seed dispersal in a tropical forest vertebrate community

Summary Interactions between a large community of vertebrate frugivore-granivores (including 7 species of large canopy birds, 19 species of rodents, 7 species of ruminants, and 6 species of monkeys), and 122 fruit species they consume, were studied for a year in a tropical rainforest in Gabon.The results show how morphological characters of fruits are involved in the choice and partitioning of the available fruit spectrum among consumer taxa. Despite an outstanding lack of specificity between fruit and consumer species, consideration of simple morphological traits of fruits reveals broad character syndromes associated with different consumer taxa. Competition between distantly related taxa that feed at the same height is far more important than has been previously supposed. The results also suggest how fruit characters could have evolved under consumer pressure as a result of consumer roles as dispersers or seed predators. Our analyses of dispersal syndromes show that fruit species partitioning occurs more between mammal taxa than between mammals and birds. There is thus a bird-monkey syndrome and a ruminant-rodent-elephant syndrome. The bird-monkey syndrome includes fruit species on which there is no pre-dispersal seed predation. These fruits (berries and drupes) are brightly colored, have a succulent pulp or arillate seeds, and no protective seed cover. The ruminant-rodent-elephant syndrome includes species for which there is pre-dispersal predation. These fruits (all drupes) are large, dull-colored, and have a dry fibrous flesh and well-protected seeds.     

Evolution of fruit types and seed dispersal:A phylogenetic and ecological snapshot

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angio-sperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats: an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

果实类型和种子传播的进化:系统发育和生态学简论

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angiosperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

Tobacco to tomatoes: a phylogenetic perspective on fruit diversity in the Solanaceae

The Solanaceae contains many species of agricultural importance. Several of these are cultivated for their fruits, such as the tomato, the pepper and the aubergine. The family is very diverse in fruit type with capsules, drupes, pyrenes, berries, and several sorts of dehiscent non-capsular fruits occurring in the 90+ genera. In this paper, recent work on fruit type evolution in angiosperms is reviewed in relation to dispersal agents and habitat ecology. Defining fruit types in the Solanaceae in a simple five state system, then mapping them onto a previously published molecular phylogeny based on chloroplast DNA allows discussion of the evolution of these fruit types in a phylogenetic framework. Capsules are plesiomorphic in the family, and although berries are a synapomorphy (shared derived character) for a large clade including the genus Solanum (tomatoes and aubergines), they have arisen several times in the family as a whole. Problems with homology of drupes and pyrenes are discussed, and areas for future investigation of fruit structure homology identified. The distribution of fruit types in the large and diverse genus Solanum is also discussed in the light of monophyletic groups identified using chloroplast gene sequences. This variety is related to recent advances in the understanding of the molecular biology of fruit development. Finally, several key areas of future comparative, phylogenetic investigation into fruit type evolution in the family are highlighted.     

Large-scale phylogenetic analyses reveal fagalean diversification promoted by the interplay of diaspores and environments in the Paleogene

The order Fagales (ca. 1325 species, 32 genera) is one of the most important orders of woody angiosperms in both tropical and temperate forests. Fagalean plants have diverse diaspores and dispersal modes, and have abundant macrofossil records. Here, we present, to our knowledge, the most comprehensive phylogenetic analysis of Fagales to date based on five plastid loci. We reconstructed the phylogenetic relationships within Fagales using parsimony, likelihood and Bayesian approaches. We inferred the evolutionary shifts of diaspore types, dispersal modes, habitats, and pollination syndromes, and estimated divergence times and rates of diversification. Fossil fruit records of Fagales were also reviewed. Our results suggest that fagalean families are all monophyletic and 29 of the 32 genera are monophyletic. The majority of inter-familial and inter-generic relationships were well resolved. Our evolutionary reconstructions indicate that winged diaspore is synapomorphic, and animal dispersal system is symplesiomorphic in Fagales. Within the order, the families diverged in the mid-Cretaceous but mainly diversified after the Cretaceous–Paleogene (K–Pg) boundary. The overwhelming majority of winged and wingless fruited genera diverged or diversified during the Paleogene, reflecting adaptation to wind and animal dispersals, respectively. Correlated evolution analyses strongly supported the correlated transitions between dispersal mode and habitat. The winged fruited groups often inhabit open habitats and are commonly dispersed abiotically, whereas the wingless fruits usually inhabit closed habitats and have a vertebrate-dispersal mode. Environmental changes triggered increased diversification of Fagales in the Paleogene together with evolution of diaspores and dispersal modes. Additionally, our paleobotanically calibrated time-scale for Fagales may be useful for ecological and physiological studies.     

Seed mass and the evolution of fleshy fruits in angiosperms

Fleshy fruits, like drupes and berries, have evolved many times through angiosperm history. Two hypotheses suggest that fleshy fruit evolution is related to changes in the seed mass fitness landscape. The reduced dispersal capability following from an increase in seed mass may be counterbalanced by evolution of traits mediating seed dispersal by animals, such as fleshy fruits. Alternatively, increasing availability and capabilities of frugivores promote evolution of fleshy fruits and allow an increase in seed size. Both these hypotheses predict an association between evolution of fleshy fruits and increasing seed size. We investigated patterns of fruit and seed evolution by contrasting seed mass between fleshy and non‐fleshy fruited sister clades. We found a consistent association between possession of fleshy fruits and heavier seeds. The direction of fruit type change did not alter this pattern; seed mass was higher in clades where fleshy fruits evolved and lower in clades where non‐fleshy fruits evolved, as compared to their sister clades. These patterns are congruent with the predictions from the two hypotheses, but other evidence is needed to distinguish between them. We emphasize the need to integrate studies of seed disperser effectiveness, seed morphology, and plant recruitment success to better understand the frugivores’ role in fleshy fruit evolution.     

Patterns in the evolution of amentiferous fruits

The diversity of fruit types within the Amentiferae appears to be the result of different seed dispersal and seedling establishment strategies that have evolved independently in the amentiferous families within the confines imposed by the wind-pollination syndrome. Dispersal of unadorned fruits and seeds by wind (e.g.,Betula, Rhoiptelea, Casuarina) has preceded the development of more efficient air-borne devices (e.g.,Ostrya, Carpinus, Engelhardia). Animal dispersal is the most advanced strategy, relying rarely on drupes, as in the shrubby Myricaceae, or on nuts, as inCorylus, Quercus, Carya, Juglans, Alfaroa, etc. The pattern of seedling establishment shows a structural-functional relationship to the seeddispersal strategy. Epigeal germination predominates in plants of open habitats. Those species with small fruits rely on wind dispersal, and their seeds have a relatively short germination time, whereas species with medium-sized fruits are often animal-dispersed with seeds that may have protracted germination times. In either case the reserve food supply for the young seedling is limited, and light is needed to spark photosynthesis. Hypogeal germination has evolved independently in several amentiferous families (e.g., Betulaceae, Fagaceae, Juglandaceae). This pattern is associated with closed habitats and plants with large seeds that have the capability of establishing vigorous seedlings in microhabitats of reduced light intensity where photosynthesis is impaired. The subterranean protection afforded the cotyledons and axillary meristems is a correlative feature that may have considerable importance in seedling survival where desiccation and predation are intense.     

西双版纳热带季节雨林145个树种繁殖体特征

分析了西双版纳热带季节雨林20公顷动态监测样地中145个树种的繁殖体特征。结果表明：雨季散布的种类最多（71.03%）,在雾凉季及干热季散布的种类相对较少;果实类型以核果,蒴果,浆果居多,黑色、黄色、褐色果实最为常见;动物散布的种类最多（56.55%）,其次为机械散布,独立散布,风力散布;大种子树种（种子最大长度〉10mm）较占优势（53.10%）;树种单粒种子重量从2.3×10^-5～22.29g不等,但76.55%的种子的重量集中在0.01～10g之间;干热季散布的种子平均重量显著高于雨季和雾凉季;独立散布的种子重量最大,风力散布的种子重量最小,而机械弹射的种子重量和动物散布的种子重量之间差异不显著;树种的成年个体最大胸径与种子重量之间存在显著正相关。     

Convergent evolution of a complex fruit structure in the tribe Brassiceae (Brassicaceae)

? Premise of study: Many angiosperms have fruit morphologies that result in seeds from the same plant having different dispersal capabilities. A prime example is found in the Brassiceae (Brassicaceae), which has many members with segmented or heteroarthrocarpic fruits. Since only 40% of the genera are heteroarthrocarpic, this tribe provides an opportunity to study the evolution of an ecologically significant novelty and its variants. ? Methods: We analyzed nuclear (PHYA) and plastid (matK) sequences from 66 accessions using maximum parsimony, maximum likelihood, and Bayesian inference approaches. The evolution of heteroarthrocarpy and its variants was evaluated using maximum parsimony and maximum likelihood ancestral state reconstructions. ? Key results: Although nuclear and plastid phylogenies are incongruent with each other, the following findings are consistent: (1) Cakile, Crambe, Vella, and Zilla lineages are monophyletic; (2) the Nigra lineage is not monophyletic; and (3) within the Cakile clade, Cakile, Didesmus, and Erucaria are paraphyletic. Despite differences in the matK and PHYA topologies at both deep and shallow nodes, similar patterns of morphological evolution emerge. Heteroarthrocarpy, a complex morphological trait, has evolved multiple times across the tribe. Moreover, there are convergent transitions in dehiscence capabilities and fruit disarticulation across the tribe. ? Conclusions: We present the first explicit analysis of fruit evolution within the Brassiceae, which exemplifies evolutionary lability. The repeated loss and gain of segment dehiscence and disarticulation suggests conservation in the genetic pathway controlling abscission with differential expression across taxa. This study provides a strong foundation for future studies of mechanisms underlying variation in dispersal capabilities of Brassiceae.     

Palynological characters and their phylogenetic signal in Rubiaceae

In the 1990s Rubiaceae became a hot spot for systematists, mainly due to the comprehensive treatment of the family by Robbrecht in 1988. Next to the exploration of macromolecular characters to infer the phylogeny, the palynology of Rubiaceae finally received the attention it deserves. This article aims to present a state-of-the-art analysis of the systematic palynology of the family. The range of varíation in pollen morphology is wide, and some of the pollen features are not known from other angiosperm taxa; e.g., a looplike or spiral pattern for the position of apertures in pantoaperturate grains. We compiled an online database at the generic level for the major pollen characters and orbicule presence in Rubiaceae. An overview of the variation is presented here and illustrated per character: dispersal unit, pollen size and shape, aperture number, position and type, sexine ornamentation, nexine pattern, and stratification of the sporoderm. The presence/absence and morphological variation of orbicules at the generic level is provided as well. The systematic usefulness of pollen morphology in Rubiaceae is discussed at the (sub)family, tribal, generic, and infraspecific levels, using up-to-date evolutionary hypotheses for the different lineages in the family. The problems and opportunities of coding pollen characters for cladistic analyses are also treated.     

The Ecology and Evolution of Fruit Odor: Implications for Primate Seed Dispersal

Primates are now known to possess a keen sense of smell that serves them in various contexts, including feeding. Many primate species are frugivorous and provide essential seed dispersal services to a variety of plants. Studies of pollination ecology, and recently seed dispersal ecology, indicate that animal mutualist behavior exerts selection pressures that drive changes in flower and fruit traits. As a result, the use of olfaction in in primate feeding ecology may have affected the evolution of fruit odor in species that rely on primate seed dispersal. However, this hypothesis is seldom tested. Here, we summarize the available information on how primates may have affected the evolution of fruit odor. We ask what the chemistry of primate fruit odor may look like, what information fruit odor may convey, whether there are geographical differences in fruit odor, and what other factors may affect the odor of fruits consumed by primates. We identify many gaps in the available data and offer research questions, hypotheses, and predictions for future studies. Finally, to facilitate standardization in the field, we discuss methodological issues in the process of odor sampling and analysis.     

Major Clades of Australasian Rutoideae (Rutaceae) Based on rbcL and atpB Sequences

Background

Rutaceae subfamily Rutoideae (46 genera, c. 660 species) is diverse in both rainforests and sclerophyll vegetation of Australasia. Australia and New Caledonia are centres of endemism with a number of genera and species distributed disjunctly between the two regions. Our aim was to generate a high-level molecular phylogeny for the Australasian Rutoideae and identify major clades as a framework for assessing morphological and biogeographic patterns and taxonomy.

Methodology/Principal Findings

Phylogenetic analyses were based on chloroplast genes, rbcL and atpB, for 108 samples (78 new here), including 38 of 46 Australasian genera. Results were integrated with those from other molecular studies to produce a supertree for Rutaceae worldwide, including 115 of 154 genera. Australasian clades are poorly matched with existing tribal classifications, and genera Philotheca and Boronia are not monophyletic. Major sclerophyll lineages in Australia belong to two separate clades, each with an early divergence between rainforest and sclerophyll taxa. Dehiscent fruits with seeds ejected at maturity (often associated with myrmecochory) are inferred as ancestral; derived states include woody capsules with winged seeds, samaras, fleshy drupes, and retention and display of seeds in dehisced fruits (the last two states adaptations to bird dispersal, with multiple origins among rainforest genera). Patterns of relationship and levels of sequence divergence in some taxa, mostly species, with bird-dispersed (Acronychia, Sarcomelicope, Halfordia and Melicope) or winged (Flindersia) seeds are consistent with recent long-distance dispersal between Australia and New Caledonia. Other deeper Australian/New Caledonian divergences, some involving ant-dispersed taxa (e.g., Neoschmidia), suggest older vicariance.

Conclusions/Significance

This comprehensive molecular phylogeny of the Australasian Rutoideae gives a broad overview of the group’s evolutionary and biogeographic history. Deficiencies of infrafamilial classifications of Rutoideae have long been recognised, and our results provide a basis for taxonomic revision and a necessary framework for more focused studies of genera and species.     

绒毛白蜡二态型果实的扩散特征

多态型果实或种子的出现对植物种群的扩散具有重要的意义。绒毛白蜡(Fraxinus velutina)的果实具有二态型特征,主要表现在果翅数量上不同,分别定义为二翅型和三翅型果实,为了比较两类果实在风传扩散时的差异,研究了两类果实的形态、果翅结构和扩散距离及扩散时长。在大型封闭地下室内,以电扇在不同速度档位产生的气流作为风源,分别从2、1.5、1m处手动释放果实,对风速为0、4.6、6.5、7.3m/s时的果实扩散距离及扩散时长进行了比较;并在此基础上对果实的形态特征与扩散特征进行了线性相关分析。结果表明:在同一高度及相同风速下,三翅型果实的水平扩散距离都极显著的大于二翅型,但其相应的扩散时长都小于二翅型。在相同情况下,三翅型的果实沉降速度显著高于二翅型。两类果实随着释放高度的增加,其扩散距离和扩散时长都相应的增加;随着风速的升高,其扩散距离及扩散时长都相应的增加。三翅型果实质量显著高于二翅型,相反,三翅型果翅长与宽都显著小于二翅型。两种翅型的果翅细胞结构都一样,细胞内部都呈现气囊状,果翅表面沿纵轴方向有流线型的纵棱。通过直线相关分析发现,翅型是对扩散距离和扩散时长影响最显著的形态特征;与果实释放高度相比,风速是影响绒毛白蜡果实扩散距离与扩散时长最显著的环境因素。绒毛白蜡三翅型果实比二翅型果实传播的远,关键在于其具有三翅,三翅对阵风瞬间响应,使得沉降速度较二翅型高,可以在瞬时风的作用下,快速传播到较远的距离。三翅型与二翅型在扩散方式上的结合增强了绒毛白蜡的生存与定殖机会。     

Frugivore gape size and the evolution of fruit size and shape in southern hemisphere floras

Abstract The present study uses differences among frugivore faunas of the southern hemisphere landmasses to test whether frugivore characteristics have influenced the evolution of fruit traits. Strong floristic similarities exist among southern landmasses; for example, 75% of New Zealand vascular plant genera also have species in Australia. However, plants in Australia and South America have evolved in the presence of a range of mammalian frugivores, whereas those in New Zealand, New Caledonia and the Pacific Islands have not. In addition, the avian frugivores in New Zealand and New Caledonia are generally smaller than those of Australia. If frugivore characteristics have influenced the evolution of fruit traits, predictable differences should exist between southern hemisphere fruits, particularly fruit size and shape. Fruit dimensions were measured for 77 New Zealand species and 31 Australian species in trans‐Tasman genera. New Zealand fruits became significantly more ellipsoid in shape with increasing size. This is consistent with frugivore gape size imposing a selective pressure on fruit ingestability. This result is not a product of phylogenetic correlates, as fruit length and width scaled isometrically for Australian species in genera shared with New Zealand. Within‐genus contrasts between New Zealand and Australian species in 20 trans‐Tasman genera showed that New Zealand species have significantly smaller fruits than their Australian counterparts. Within‐genus contrasts between New Zealand and South American species in nine genera gave the same result; New Zealand species had significantly smaller fruits than their South American counterparts. No difference was found in fruit size or shape between New Zealand and New Caledonia congeneric species from 12 genera. These results are consistent with the broad characteristics of the frugivore assemblage influencing the evolution of fruit size and shape in related species. The smaller‐sized New Zealand frugivore assemblage has apparently influenced the evolution of fruit size of colonizing taxa sometimes within a relatively short evolutionary timeframe.     

Associated evolution of fruit size,fruit colour and spines in Neotropical palms

Understanding how ecological interactions have shaped the evolutionary dynamics of species traits remains a challenge in evolutionary ecology. Combining trait evolution models and phylogenies, we analysed the evolution of characters associated with seed dispersal (fruit size and colour) and herbivory (spines) in Neotropical palms to infer the role of these opposing animal–plant interactions in driving evolutionary patterns. We found that the evolution of fruit colour and fruit size was associated in Neotropical palms, supporting the adaptive interpretation of seed‐dispersal syndromes and highlighting the role of frugivores in shaping plant evolution. Furthermore, we revealed a positive association between fruit size and the presence of spines on palm leaves, bracteas and stems. We hypothesize that interactions between palms and large‐bodied frugivores/herbivores may explain the evolutionary relationship between fruit size and spines. Large‐bodied frugivores, such as extinct megafauna, besides consuming the fruits and dispersing large seeds, may also have consumed the leaves or damaged the plants, thus simultaneously favouring the evolution of large fruits and defensive structures. Our findings show how current trait patterns can be understood as the result of the interplay between antagonistic and mutualistic interactions that have happened throughout the evolutionary history of a clade.     

Gynoecium, fruit and seed structure of Paullinieae (Sapindaceae)

Despite an emphasis on fruit characters in Paullinieae taxonomy, few detailed morphological and anatomical studies of the gynoecia, fruits and seeds exist. The aims of the present study were (1) to provide a detailed documentation of gynoecium, fruit and seed structure and ontogeny in selected Paullinieae taxa; (2) to determine whether the gynoecium, seed and seedling provide additional characters of systematic value within the tribe; and (3) to relate the structural findings to mechanisms of fruit dehiscence and dispersal within these taxa. Newly described characters of systematic value within Paullinieae are shape and surface of the obturator, type of pollen tube transmitting tract, indumentum of the inner and outer surface of the carpels, ovary wall anatomy, aril anatomy, pseudo-hilum form, seedling germination mode and structure of first leaves. The fruits of Paullinia are septifragal, and conspicuous colour contrasts between the pericarp, aril and seed in most species of this genus are suggestive of a bird dispersal syndrome. Interestingly, it appears that relatively minor structural changes are associated with switches to rodent dispersal in Paullinia sphaerocarpa and water dispersal in P. clathrata and P. hystrix. Anemochorous fruits are septifragal ( Cardiospermum and Urvillea ) or schizocarpic ( Houssayanthus , Lophostigma , Serjania ). They are structurally similar and Cardiospermum with septifragal capsules may also show septicidal dehiscence.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 159–189.     

Seed Size and Dispersal Systems of Early Cretaceous Angiosperms from Famalicão, Portugal

Seeds and fruits of Early Cretaceous (Barremian-Aptian) angiosperms from the Famalic?o locality in Portugal were analyzed to establish seed and fruit size (volume) distributions and to infer the proportion of animal-dispersed fruits. On the basis of a sample of 106 angiosperm fruit and seed taxa, the average seed size was 0.78 mm3 (range 0.02-6.86 mm3), whereas the average fruit size was 2.06 mm3 (range 0.12-8.34 mm3). Variation in seed size among taxa is smaller than in modern plant communities, but within-taxon variation is similar to that known for extant plants. No significant difference in the size of "fleshy" versus other fruits was observed. The proportion of fleshy fruits was 24.5%. This high figure was surprising and indicates that the significance of animal dispersal during an early stage in angiosperm evolution has been underestimated. We suggest that reptiles and multituberculates, and perhaps other mammals and birds as well, were the likely seed dispersers and that the early angiosperms from Famalic?o probably were herbs or small shrubs that inhabited a semiopen coniferous woodland.     

Incidence of dioecy in relation to growth form,pollination and dispersal

Summary Recent theories predict the evolution of dioecy among higher plants, in association with certain pollination and fruit dispersal traits. However, reported associations of dioecy with pollination, dispersal and growth form traits have not distinguished the effects of each trait separately, controlling for the others. Because these traits are associated among themselves, existing analyses may involve spurious or indirect correlations. This paper reports the incidence of dioecy in a subarctic and an arctic flora, and analyzes the occurrence of dioecy among vascular plants classified jointly by growth form, floral (pollination) syndrome, and fruit (dispersal) syndrome. Dioecism is no more frequent in the arctic flora as a whole, but its incidence increases northward among woody plants. This increase is associated with an increase in the proportion of woody species having small, inconspicuous flowers, and not with the syndrome of fleshy or animal dispersed fruits. Within the floras of Alaska, California, and the Northeastern US, dioecy is markedly more frequent among woody plants and among plants having small, inconspicuous flowers, and that is the only strong statistical association of dioecy for the species of these floras. When genera and families are analyzed similarly, dioecy is also associated significantly with dispersal syndrome. Thus, among angiosperms, evidence currently does not support either an uniquely strong or exclusive association of dioecy with dispersal traits, as it does for gymnosperms (Givnish 1980). It is extremely desirable to analyze the occurrence of dioecy among taxa classified jointly by all relevant ecological traits, rather than analyzing marginal distributions.     

PHYLOGENY OF THE RUBIACEAE AND THE LOGANIACEAE: CONGRUENCE OR CONFLICT BETWEEN MORPHOLOGICAL AND MOLECULAR DATA?

Phylogenetic analyses of 33 genera of Rubiaceae were performed using morphological and a few chemical characters. Parsimony analysis based on 29 characters resulted in eight equally parsimonious trees, with a consistency index of 0.40 and a retention index of 0.69. These results were compared to a phylogenetic analysis of the same genera based on chloroplast DNA restriction site data. There are discrepancies between the two analyses, but if we consider groupings reflected in the present classification there is much congruency. With the exception of four genera, all the genera are positioned in the same group of taxa in the two analyses. Clades of taxa representing three of the four subfamilies (~the Antirheoideae, ~the Rubioideae, and the ~Ixoroideae) are monophyletic, while the fourth subfamily Cinchonoideae is shown to be paraphyletic. Both analyses support a widened tribe Chiococceae, including the former subtribe Portlandiinae (Condamineeae). Furthermore, in both analyses the tribe Hamelieae is placed outside the subfamily Rubioideae where it is now housed. In search for the most plausible sister group to the Rubiaceae, the genus Cinchona (Rubiaceae) was analyzed together with 13 genera of the Loganiaceae, Nerium (Apocynaceae), and Exacum (Gentianaceae). Cornus (Comaceae), Olea (Oleaceae), and these two genera together were used as outgroups. The analysis, including 25 characters, 16 taxa, and with Cornus and Olea together as an outgroup, resulted in four equally parsimonious trees, with a consistency index of 0.53 and a retention index of 0.62. The non-Loganiaceae taxa Cinchona (Rubiaceae), Nerium (Apocynaceae), and Exacum (Gentianaceae) were all found to have their closest relatives within the Loganiaceae indicating that the Loganiaceae are paraphyletic and ought to be reclassified. As a result of the morphological data the most plausible sister group to the Rubiaceae is the tribe Gelsemieae of the Loganiaceae.     

Cryptic clades, fruit wall morphology and biology of Andira (Leguminosae: Papilionoideae)

Andira comprises 29 species distributed throughout tropical America, with two subspecies in Africa. Its fruits are unusual for a papilionoid legume because they are drupes. The majority of species have fruits dispersed by bats, but eight species have larger fruits dispersed by-rodents. Some fruits of both dispersal types are secondarily dispersed by water. Cladistic analysis of chloroplast DNA (cpDNA) restriction site characters discovered four well-supported clades of Andira species. None of these 'cryptic' clades had been recognized by previous workers, because they are not apparently marked by any known morphological innovations. This prompted a search for new characters that might support these groupings. An anatomical study of fruit walls of 25 Andira species revealed the presence of three principal endocarp types, dominated by (1) parenchyma, (2) fibres, or (3) stone cells. These features arc best coded as a single unordered multistate character. When incorporated into a simultaneous cladistic analysis of previously gathered molecular and morphological data, states of this endocarp character are shown to be apomorphies for two of the well-supported clades evident in the cpDNA restriction site data. The most likely plesiomorphic state for the endocarp is parenchyma-dominated. Thicker, stronger endocarps of fibres and stone cells may have evolved in response to the need to protect the seed from predators.